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Adapting defensive behavior to the characteristics of a threatening situation is a fundamental function of the brain. Particularly, threat imminence plays a major role for the organization of defensive responses. Acute threat prompts phasic physiological responses, which are usually associated with an intense feeling of fear. In contrast, diffuse and potentially threatening situations elicit a sustained state of anxious apprehension. Detection of the threatening stimulus defines the key event in this framework, initiating the transition from potential to acute threat. Consequently, attention to threat is crucial for supporting defensive behavior. The functions of attention are finely tuned to the characteristics of a threatening situation. Potential threat is associated with hypervigilance, in order to facilitate threat detection. Once a threatening stimulus has been identified, attention is selectively focused on the source of danger. Even though the concepts of selective attention and hypervigilance to threat are well established, evidence for their neural correlates remain scarce. Therefore, a major goal of this thesis is to elucidate the neural correlates of selective attention to acute threat and hypervigilance during potential threat. A second aim of this thesis is to provide a mechanistic account for the interaction of fear and anxiety. While contemporary models view fear and anxiety as mutually exclusive, recent findings for the neural networks of fear and anxiety suggest potential interactions. In four studies, aversive cue conditioning was used to induce acute threat, while context conditioning served as a laboratory model of potential threat. To quantify neural correlates of selective attention and hypervigilance, steady-state visual evoked potentials (ssVEPs) were measured as an index of visuocortical responding. Study 1 compared visuocortical responses to acute and potential threat for high versus low trait-anxious individuals. All individuals demonstrated enhanced electrocortical responses to the central cue in the acute threat condition, suggesting evidence for the neural correlate of selective attention. However, only low anxious individuals revealed facilitated processing of the contexts in the potential threat condition, reflecting a neural correlate of hypervigilance. High anxious individuals did not discriminate among contexts. These findings contribute to the notion of aberrational processing of potential threat for high anxious individuals. Study 2 and 3 realized orthogonal combinations of cue and context conditioning to investigate potential interactions of fear and anxiety. In contrast to Study 1 and 2, Study 3 used verbal instructions to induce potentially threatening contexts. Besides ssVEPs, threat ratings and skin conductance responses (SCRs) were recorded as efferent indices of defensive responding. None of these studies found further evidence for the neural correlates of hypervigilance and selective attention. However, results for ratings and SCRs revealed additive effects of fear and anxiety, suggesting that fear and anxiety are not mutually exclusive, but interact linearly to organize and facilitate defensive behavior. Study 4 tested ssVEPs to more ecologically valid forms of context conditioning, using flickering video stimuli of virtual offices to establish context representations. Contrary to expectations, results revealed decreased visuocortical responses during sustained presentations of anxiety compared to neutral contexts. A disruption of ssVEP signals eventually suggests interferences by continuously changing video streams which are enhanced as a function of motivational relevance. In summary, this thesis provided evidence for the neural correlates of attention only for isolated forms of fear and anxiety, but not for their interaction. In contrast, an additive interaction model of fear and anxiety for measures of defensive responding offers a new perspective on the topography of defensive behavior.
Abstract
Tobacco addiction is considered as a chronic relapsing disorder, characterized by compul-sive drug seeking and intake. Learning processes are stressed to account for the situational-specific expression of core features of the disorder, e.g., craving for drug, tolerance and ex-cessive consumption. According to incentive theories, smoke conditioned stimuli are hy-pothesized to be appetitive in nature, promoting craving, approach and consummatory be-havior. Commonly, smoking cues are treated as simple excitatory conditioned stimuli formed by a close and reliable overlap with the drug effect. However, the smoking ritual comprises a multitude of stimuli which may give rise to different forms of learning and con-ditioned responses partially opposing each other. Previous research suggests the predictive content and the temporal proximity of smoking stimuli to the drug effect as important de-terminants of cue reactivity. In contrast to stimuli related to the preparatory stage of smok-ing and the start of consumption (BEGIN stimuli), stimuli from the terminal stage of smok-ing (END stimuli) apparently lack high cue reactivity. Several lines of evidence suggest the poor cue properties of terminal stimuli to be related to their signaling of poor smoke availa-bility. Indeed, cue reactivity is commonly decreased when smoking appears to be unavaila-ble. Moreover, the learning literature suggests that stimuli predictive for the non-availability of reward may acquire the capacity to modulate or oppose the responses of ex-citatory conditioned stimuli. Therefore, the aim of the present thesis was to enhance our knowledge of stimulus control in human drug addiction and incentive motivation by running a series of conditioning studies with smoke intake and monetary reward as reinforcer. Sub-jective report and physiological measures of motivational valence and consummatory re-sponse tendencies were used as dependent variables.
The first experiment of this thesis used a differential conditioning paradigm to reveal evi-dence for the conditioning of preparatory and consummatory responses to a CS+ for smok-ing. Neutral pictograms served as CSs and single puffs on a cigarette as US. In line with the predictions of incentive theories, the excitatory CS+ for smoking acquired the ability to evoke an appetitive conditioned response, as indicated by enhanced activity of the M. zy-gomaticus major. Moreover, anticipation of puffing on the cigarette increased the activity of the M. orbicularis oris (lip muscle), indicating the activation of consummatory response tendencies. Finally, the CS+ evoked stronger skin conductance responses, indicative of in-creased autonomic arousal and orienting in preparation for action. In contrast, the rating data were apparently unaffected by the experimental contingency. In sum, the physiological data provide support for the notion that excitatory smoke conditioning gives rise to appeti-tive and consummatory conditioned responses, which may at least partially contribute to the maintenance of tobacco addiction.
The second experiment of this thesis adapted the conditioning protocol of the first study to probe the functional significance of terminal stimuli in the control of addictive behavior. This study manipulated the predictive relationship of BEGIN and END stimuli to smoke rein-forcement to provide further support for the differential reactivity to both stimuli and the retarded (i.e., delayed) conditioning of END stimuli. Overall, the results of the first study of this thesis were conceptually replicated as the association of a BEGIN stimulus with smoke intake resulted in the acquisition of appetitive and consummatory physiological responses. Importantly, the results revealed evidence for a retarded excitatory conditioning of END stimuli. Thus, pairing of an END stimulus with smoke intake failed to produce a conditioned discrimination in terms of motivational valence and autonomic arousal, as indicated by the activity of the M. corrugator supercilii and the skin conductance data. These results provide further support for the notion that END stimuli may be weak cues for smoking. Moreover, in light of the results of the first study of this thesis, the retarded excitatory conditioning of terminal stimuli may be suggestive of an inhibitory response component, which may be re-lated to their signaling of poor smoke availability. In sum, these results add to a growing body of data, which suggest that the expression of cue reactivity may be modulated by the temporal proximity and the availability of the drug effect.
The aim of the third study of this thesis was to provide “proof of concept” for an inhibi-tory conditioning notion of terminal stimuli. In this analog study BEGIN and END stimuli were emulated as discriminative SD and S for monetary reward. During an acquisition phase conditioned inhibition was established to the S predictive of the non-availability of re-ward. Subsequently a retardation test was used to substantiate conditioned inhibition. In this test, excitatory conditioning of the previous S was compared to the excitatory condi-tioning of a novel control stimulus. Importantly, the results revealed evidence for reward conditioned inhibition as indicated by the retarded acquisition of subjective (pleasure and reward expectancy) and physiological (skin conductance and activity of the M. orbicularis oculi) responses. In sum, these results provide support for the notion that stimuli predictive for the non-availability of reward may acquire the capacity to oppose the responses of ex-citatory conditioned stimuli. Thus, future research may benefit from the consideration of inhibitory conditioning processes in drug addiction, which may be of theoretical, methodo-logical and clinical importance.
In sum, the present thesis revealed evidence for 1) an appetitive nature of excitatory condi-tioned smoking cues, 2) the dependency of this learning process on the temporal position of the conditioned stimuli in the intake ritual and 3) the acquisition of conditioned inhibition by a stimulus predictive for the non-availability of reward, as evident in retarded excitatory conditioning. Overall, these studies made a novel contribution to the field of human drug addiction and incentive motivation and provided valuable suggestions for further research.
In order to survive, organisms avoid threats and seek rewards. Classical conditioning is a simple model to explain how animals and humans learn associations between events that allow them to predict threats and rewards efficiently. In the classical conditioning paradigm, a neutral stimulus is paired with a biologically significant event (the unconditioned stimulus – US). In virtue of this association, the neutral stimulus acquires affective motivational properties, and becomes a conditioned stimulus (CS+). Defensive responses emerge for pairings with an aversive US (e.g., pain), and appetitive responses emerge for pairing with an appetitive event (e.g., reward). It has been observed that animals avoid a CS+ when it precedes an aversive US during a training phase (CS+ US; forward conditioning); whereas they approach a CS+ when it follows an aversive US during the training phase (US CS+; backward conditioning). These findings indicate that the CS+ acquires aversive properties after a forward conditioning, whereas acquires appetitive properties after a backward conditioning. It is thus of interest whether event timing also modulates conditioned responses in such an opponent fashion in humans, who are capable of explicit cognition about the associations. For this purpose, four experiments were conducted in which a discriminative conditioning was applied in groups of participants that only differed in the temporal sequence between CS+ onset and US onset (i.e., the interstimulus interval – ISI). During the acquisition phase (conditioning), two simple geometrical shapes were presented as conditioned stimuli. One shape (CS+) was always associated with a mild painful electric shock (i.e., the aversive US) and the other one (CS-) was never associated with the shock. In a between-subjects design, participants underwent either forward or backward conditioning. During the test phase (extinction), emotional responses to CS+ and CS- were tested and the US was never presented. Additionally, a novel neutral shape (NEW) was presented as control stimulus. To assess cognitive components, participants had to rate both the valence (the degree of unpleasantness or pleasantness) and the arousal (the degree of calmness or excitation) associated with the shapes before and after conditioning. In the first study, startle responses, an ancestral defensive reflex consisting of a fast twitch of facial and body muscles evoked by sudden and intense stimuli, was measured as an index of stimulus implicit valence. Startle amplitude was potentiated in the presence of the forward CS+ whilst attenuated in the presence of the backward CS+. Respectively, the former response indicates an implicit negative valence of the CS+ and an activation of the defensive system; the latter indicated an implicit positive valence of the CS+ and an activation of the appetitive system. In the second study, the blood-oxygen level dependent (BOLD) response was measured by means of functional magnetic resonance imaging (fMRI) to investigate neural responses after event learning. Stronger amygdala activation in response to forward CS+ and stronger striatum activation in response to backward CS+ were found in comparison to CS-. These results support the notion that the defensive motivational system is activated after forward conditioning since the amygdala plays a crucial role in fear acquisition and expression. Whilst the appetitive motivational system is activated after backward conditioning since the striatum plays a crucial role in reward processing. In the third study, attentional processes underlying event learning were observed by means of steady-state visual evoked potentials (ssVEPs). This study showed that both forward and backward CS+ caught attentional resources. More specifically, ssVEP amplitude was higher during the last seconds of forward CS+ that is just before the US, but during the first seconds of backward CS+ that is just after the US. Supposedly, attentional processes were located at the most informative part of CS+ in respect to the US. Participants of all three studies rated both forward and backward CS+ more negative and arousing compared to the CS-. This indicated that event timing did not influence verbal reports similarly as the neural and behavioral responses indicating a dissociation between the explicit and implicit responses. Accordingly, dual process theories propose that human behavior is determined by the output of two systems: (1) an impulsive implicit system that works on associative principles, and (2) a reflective explicit system that functions on the basis of knowledge about facts and values. Most importantly, these two systems can operate in a synergic or antagonistic fashion. Hence, the three studies of this thesis congruently suggest that the impulsive and the reflective systems act after backward association in an antagonistic fashion. In sum, event timing may turn punishment into reward in humans even though they subjectively rate the stimulus associated with aversive events as being aversive. This dissociation might contribute to understand psychiatric disorders, like anxiety disorders or drug addiction.